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ACTA AMAZONICA http://dx.doi.org/10.1590/1809-4392201700602

ORIGINAL ARTICLE Silvicultural performance of five forest species in the central Brazilian Amazon Murilo Rezende MACHADO1*, Rodrigo CAMARA2, Paulo de Tarso Barbosa SAMPAIO 3, João Baptista Silva FERRAZ 3, Marcos Gervasio PEREIRA4 1 Instituto Chico Mendes de Conservação da Biodiversidade, Floresta Nacional do Rio Preto, Zona Rural, Conceição da Barra, 2 Universidade Federal Rural do Rio de Janeiro, Programa de Pós Graduação em Ciências Ambientais e Florestais, Programa Nacional de Pós Doutorado da Coordenação de Aperfeiçoamento de Pessoal de Nível Superior, BR 465, Km 7, Seropédica, Brazil 3 Instituto Nacional de Pesquisas da Amazônia, Departamento de Silvicultura Tropical, Avenida André Araújo, 2936, Manaus, Brazil 4 Universidade Federal Rural do Rio de Janeiro, Departamento de Solos, BR 465, Km 7, Seropédica, Brazil * Corresponding author: [email protected]

ABSTRACT Planting of forest species of timber interest helps to reduce the deforestation pressure on the Amazon forest, promotes sustainable development of the producing region and generates ecological benefits. The objective of this work was to evaluate the survival rate and growth of four native ( macrophylla, Parkia decussata, odorata and ) and one exotic (Acacia mangium) species in monospecific (spacing of 2 x 2 m) established on areas previously used for grazing, in Itacoatiara, State of Amazonas, Brazil. When the were four years old, we collected biometric data 2 3 [height at 1.30 m (DBH, cm), crown projection area (CPA, m ), total height (Ht, m), commercial cylinder volume (Vcyl, m ha-1)], and qualitative data from visual diagnosis [survival rate (S, %), nutritional status (NS, G = good, D = deficient, %), and phytosanitary status (PS, S = satisfactory, N = non-satisfactory, %)]. Three plots of 128 2m , with 32 each, were evaluated for each species. Jacaranda copaia, followed by and Parkia decussata, were the recommended species for planting in areas with edaphoclimatic conditions similar to those of the present work, due to their better performance according to most of the variables. KEYWORDS: dendrometric variables, monospecific plantings, silviculture, growth Desempenho silvicultural de cinco espécies florestais na Amazônia central brasileira RESUMO Plantios de espécies florestais de interesse madeireiro contribuem para diminuir a pressão sobre a floresta amazônica, proporcionam o desenvolvimento sustentável na região e geram benefícios ecológicos. Este estudo objetivou avaliar a sobrevivência e o crescimento de quatro espécies nativas (, Parkia decussata, Dipteryx odorata e Jacaranda copaia) e uma exótica (Acacia mangium), em plantios monoespecíficos (espaçamento de 2 x 2 m) estabelecidos em áreas anteriormente ocupadas por pastagem, em Itacoatiara, Amazonas. Aos quatro anos de idade, foram obtidos dados biométricos 2 [diâmetro à altura de 1,30 m do solo (DBH, cm), área de projeção de copa (CPA, m ), altura total (Ht, m), volume comercial do 3 -1 cilindro (Vcyl, m ha )], e dados qualitativos obtidos por diagnose visual [taxa de sobrevivência (S; %), estado nutricional (NS, G: bom; D: deficiente; %) e estado fitossanitário (PS, S: satisfatório; NS: não-satisfatório; %). Para cada espécie considerou- se três parcelas de 128 m2, cada uma com 32 plantas. Jacaranda copaia, seguida de Dipteryx odorata e Parkia decussata, foram as espécies mais recomendadas para o plantio em áreas com condições edafoclimáticas semelhantes às do presente trabalho, devido ao seu melhor desempenho para a maioria das variáveis. PALAVRAS-CHAVE: variáveis dendrométricas, plantios monoespecíficos, silvicultura, crescimento de árvores

CITE AS: Machado, M.R.; Camara, R.; Sampaio, P.T.B.; Ferraz, J.B.S.; Pereira, M.G. 2018. Silvicultural performance of five forest species in the central Brazilian Amazon. Acta Amazonica 48: 10-17.

10 VOL. 48(1) 2018: 10 - 17 ACTA AMAZONICA MACHADO et al. Silvicultural performance of forest species

INTRODUCTION for cattle grazing purposes. In December 2003, the owners of the farm isolated part of the pasture (36 ha) in order to install The Brazilian Legal Amazon, whith an area of six million commercial monospecific plantations of five forest species, by kilometers (approximately 60% of the entire national territory), means of seedlings. The planted trees were spaced 2 x 2 m apart, is known for its rich natural resources. However, estimates and four native Amazonian forest species were selected: Dipteryx indicate that 20% of the total original forest of the region has odorata (Aubl.) Willd. (, non-pioneer, planted area already been deforested (IBGE, 2010), in order to serve the of 0.80 ha), Jacaranda copaia (Aubl.) D Don (, interests of the timber industry, among others (Diniz et al. 2009). pioneer, 5.20 ha), Parkia decussata Ducke (Mimosaceae, non- The situation is currently of major global environmental concern pioneer, 2.76 ha), and Swietenia macrophylla King. (, (Vieira et al. 2008). In parallel, the Brazilian participation in non-pioneer, 1.60 ha). Additionally, an exotic species originated the world trade of forest products has been increasing (Silva et from Oceania, Acacia mangium Willd. (Fabaceae, pioneer, al. 2013a), with a predominant participation of the northern 7.50 ha), was also selected. Machado et al. (2016) provide states. Large areas of extensive livestock farming have been detailed information on the preparation of the area prior to replaced by forest plantations with native and exotic species, in the establishment of the plantations. order to supply the demand for forest products and to comply Acacia mangium, Dipteryx odorata and Swietenia with environmental legislation (ABRAF 2013). macrophylla are considered suitable for cultivation in The above scenario makes reforestation programs a priority Amazonian edaphoclimatic conditions because they show to achieve both the decrease of deforestation pressure on high retranslocation rates of phosphorous and nitrogen, which the remaining area of primary Amazon forests, maintaining are the most limiting nutrients regarding the low availability to environmental services such as erosion control, carbon plants in tropical soils (Machado et al. 2016). According to the sequestration, hidrological balance, biodiversity conservation, latter authors, Parkia decussata and Jacaranda copaia both have and the increase of income for rural properties in the region high retranslocation rates of phosphorous. In monospecific (Galeão et al. 2006; Tonini et al. 2006). However, the plantations, Swietenia macrophylla is able to enrich the soil achievement of these ecological benefits depends on improved availability of potassium and phosphorous, while the soil under information on silvicultural aspects of candidate species for Jacaranda copaia presents high values of calcium, sum of bases programs, such as stem shape, growth, resistance and pH, up to the depth of 0.30 m (Machado et al. 2017). In to pests and nutritional and edaphic requirements, allowing rural properties in the Brazilian Amazon region, Acacia mangium the selection of the more appropriate species (Nascimento et has been planted for the production of for export (Barros al. 2012). Unfortunately, the information on the silvicultural et al. 2009), whereas Jacaranda copaia has been used to provide performance of many exotic species and, especially, native species thermal comfort for cattle due to shading, and also for the future in Amazonian conditions is still incipient (Costa et al. 2015). obtaining of wood, and Dipteryx odorata has been planted for In this context, the objective of this work was to evaluate medicinal purposes (Santos and Mitja 2010). and the silvicultural performance of four native (Swietenia of Dipteryx odorata are used in the form of tea and also with macrophylla, Parkia decussata, Dipteryx odorata and Jacaranda alcohol maceration, for internal use, to combat headaches, copaia) and one exotic (Acacia mangium) species, in cough, indigestion, and epilepsy (Vázquez et al. 2014). monospecific plantations established on areas previously used In December 2006, we established three plots of 128 m2, for grazing, in Itacoatiara, State of Amazonas, Brazil. with 32 trees each, per species plantation. The criteria used to locate the plots were the standardization of the size of the MATERIAL AND METHODS planted areas by species, and similar relief conditions (plan). We considered a completely randomized experimental design The experiment was conducted in the Nova Vida farm (230 with five treatments (species) and three replications (plots) for ha), located along the State Highway AM-010, km 250, in each species. Initially the following variables were measured: Itacoatiara, State of Amazonas (AM), Brazil (2º56’13”S, (a) diameter at breast height (1.30 m) (DBH, cm), measured 58º55’56”W). The climate of the region is type Am (Tropical with a diametrical tape; (b) total height (Ht, m), measured monsoon), very rainy (≥ 2500 mm per year), with dry winters with a graduated rod; and (c) commercial cylinder volume (Reboita et al. 2010). The relief of the plantation area varied 3 -1 (Vcyl, m ha ). The cylV was obtained by the equation Vcyl = G from flat to hilly, and the predominant soil type was classified 2 × Ht × FF, in which G is the basal area per hectare (m ), Ht is as Oxisoil (Falesi et al. 1969). the mean total height (m) and FF is the average commercial In the 1990’s, trees of high economic value were initially form factor for the species. The FF value was obtained by the removed from the original terra-firme dense ombrophylous equation FF = OV / CV, in which OV is the observed volume forest that covered the area. Afterwards, the remaining (rigorous scaling) and CV is the cylinder volume, considering vegetation was felled and burned, and the area was sowed DBH and the commercial height of the tree. The trees were with the grass Brachiaria humidicula (Rendle) Schweickerdt standing on the ground during this procedure, which is called

11 VOL. 48(1) 2018: 10 - 17 ACTA AMAZONICA MACHADO et al. Silvicultural performance of forest species the Smalian method (Soares et al. 2011). Thus, the volume of et al. (2017), and excluded the variables or soil attributes that the stems (with bark) was determined by successive application had low correlation (< 0.70%) with the principal axis 1 or axis of the Smalian method on sections 1.0, 2.0 and 3.0 m long 2. Multivariate analyses were carried out using version 2.17c with a diameter limit of approximately 3 cm, using a diametric of the PAST program (Hammer et al. 2001). tape and ruler tape. For this purpose, we scaled two trees of average DBH for each species and plot. The use of a specific RESULTS FF for each species and site is very important to evaluate The species presented the following descending order of volumetric production. Although this factor may contribute number of stems (units per hectare): Acacia mangium (5,138) to underestimate the commercial volume (Tonini et al. 2006), > Dipteryx odorata (3,020) > Swietenia macrophylla (2,915) > comparisons between plantings of various species at different Parkia decussata (2,599) > Jacaranda copaia (2,495). Jacaranda sites are more accurate when using this factor. copaia had low rates of stem bifurcation and higher rates of In December 2007, we measured: (a) survival rate (S, 2 both rectilinear stem and circular crown (Table 1). Individuals %); (b) crown projection area (CPA, m ), measured with a of this species, as well as of Dipteryx odorata and Parkia measuring tape. CPA was calculated through the ellipse area decussata, had the highest rates of satisfactory phytosanitary equation CPA = P × P , in which P is the crown projection bl dl bl status and good nutritional status (Table 1). between planting lines, and Pdl is the crown projection in the direction of planting line (π = 3, 14). All initial variables At four years of age, survival rates of all species exceeded (DBH, H , and V ) were also measured again. We estimated 80%, with the exception of Jacaranda copaia, which presented t cyl a survival rate < 70% (Table 1). We observed significant the mean annual increment per area for height (MAI-Ht, m -1 -1 differences among the species in DBH (F = 104.506, p < yr ), diameter (MAI-DBH, cm yr ) and volume (MAI-Vcyl, m3 ha-1 yr-1), and also the current annual increment per area 0.001) (Table 2). Jacaranda copaia presented a significantly -1 -1 higher mean DBH than the other species, followed by for height (CAI-Ht, m yr ), diameter (CAI-DBH, cm yr ) 3 -1 -1 Parkia decussata. Acacia mangium and Swietenia macrophylla and volume (CAI-Vcyl, m ha yr ), by comparing the results obtained in 2006 and 2007. presented intermediate mean DBH, while Dipteryx odorata presented the lowest value. This pattern was confirmed by the We also estimated qualitative variables by visual inspection MAI-DBH and CAI-DBH results, which, in general, tended of the trees (Galeão et al. 2006): (a) stem bifurcation (B, %); to be higher for Jacaranda copaia and Parkia decussata, and (b) stem quality (LQ, R = rectilinear, T = tortuous); (c) crown Dipteryx odorata quality (CQ, C = circular, I = irregular); (d) nutritional status lower for (Table 3). (NS, B = good, D = deficient); and (e) phytosanitary status Total height varied significantly among the species (F (PS, S = satisfactory, NS = non-satisfactory). = 186.229, p < 0.001) (Table 2). Acacia mangium had the highest mean for this variable, followed by Jacaranda copaia. DBH, CPA, Ht and Vcyl data were subjected to analysis of variance, followed by the LSD test to compare the means (5% Dipteryx odorata, Parkia decussata, and Swietenia macrophylla significance level). Hierarchical cluster analysis (Ward’s method) had the lowest values in decreasing order. This pattern was also provided a dendrogram for the classification of forest species reflected in MAI-Ht and CAI-Ht, which had higher values for Jacaranda copaia and Parkia decussata, and lower values for performance, using the mean values of DBH, CPA, Ht, Vcyl, S, NS-B, PS-S, B, LQ-R for each species. Principal component Swietenia macrophylla (Table 3). analysis was performed with the purpose of identifying the The mean CPA differed significantly among the five species biometric variables that correlated with the monospecific (F = 7.303, p < 0.001) (Table 2). Dipteryx odorata, Jacaranda plantations. In this latter analysis, we also considered the mean copaia and Parkia decussata had higher mean values of CPA, values of soil chemical attributes that were provided by Machado and did not differ significantly among each other.Acacia

Table 1. Survival rate (S), stem bifurcation (B), stem quality (SQ, R = rectilinear, T = tortuous), crown quality (CQ, C = circular, I = irregular), phytosanitary status (PS, S = satisfactory, N = non-satisfactory) and nutritional status (NS, G = good, D = deficient) of five forest species, in 4-year-old homogeneous plantations in Itacoatiara, State of Amazonas, Brazil. Numbers are averages of three replicates. Each replicate was a 128 m2 plot planted with 32 seedlings.

SQ (%) CQ (%) PS (%) NS (%) Species S (%) B (%) R T C I S N G D Acacia mangium 81 69 6 94 84 16 86 14 87 13 Dipteryx odorata 99 17 75 25 90 10 100 0 100 0 Jacaranda copaia 68 1 100 0 100 0 100 0 100 0 Parkia decussata 95 3 78 22 83 17 100 0 100 0 Swietenia macrophylla 98 15 71 29 95 5 0 100 71 29

12 VOL. 48(1) 2018: 10 - 17 ACTA AMAZONICA MACHADO et al. Silvicultural performance of forest species mangium and Swietenia macrophylla had the lower values and The multivariate hierarchical cluster analysis showed that did not differ significantly between them. Dipteryx odorata and Parkia decussata were very similar, with Regarding volumetry, significant differences were observed a lower binding distance, and formed a group with a slightly lower similarity (approximately double the previous binding in the mean Vcyl among the five species (F = 111.933; p < distance) with Jacaranda copaia (Figure 1). Thus, these three 0.000). Acacia mangium had a 222% higher Vcyl than the species presented higher similarity in relation to the diametric second highest value of Jacaranda copaia (Table 4). Dipteryx variables (DBH, Ht, CPA, and Vcyl). On the other hand, odorata and Swietenia macrophylla had the lower Vcyl values Dipteryx odorata, Jacaranda copaia and Parkia decussata had (Table 4). MAI-Vcyl and CAI-Vcyl followed the same pattern lower similarity with the other native species (Swietenia of Vcyl, resulting in highest values for Acacia mangium and lowest for Dipteryx odorata (Table 4). macrophylla). The exotic Acacia mangium presented the lowest similarity to the other species studied. We registered an intense attack of Swietenia macrophylla Table 2. Mean diameter at 1.30 m (DBH), total height (Ht) and crown projection area (CPA) of five forest species, in 4-year-old homogeneous plantations in (100% of the individuals) by Hypsipylla grandella (Zeller), Itacoatiara, State of Amazonas, Brazil. Numbers are averages of three replicates. Each which caused the death of older shoots and a super growth replicate was a 128 m2 plot planted with 32 seedlings. Different letters following of lateral buds. the numbers in a column indicate significant differences by the LSD test (p < 0.05). Principal component analysis grouped the plantations of 2 Dipteryx odorata and Jacaranda copaia in the right portion Species DBH (cm) Ht (m) CPA (m ) Acacia mangium 8.5±1.9 c 13.3±1.9 a 1.6±1.0 b of the axis 1 (negative values), while the other plantations Dipteryx odorata 5.2±1.1 e 7.1±1.3 c 4.6±1.9 a (Acacia mangium, Parkia decussata, and Swietenia macrophylla) were grouped in the left portion of the axis 1 (Figure 2). The Jacaranda copaia 10.9±2.7 a 11.3±2.4 b 4.0±0.6 a axis 1 explained 70.34% of the variability of the results. Axis Parkia decussata 9.8±1.6 b 6.6±1.0 c 4.4±2.2 a 2 explained 23.93% of the data variability, and located the Swietenia macrophylla 7.1±1.4 d 5.4±0.9 d 1.7±0.5 b plantations of Dipteryx odorata, Jacaranda copaia and Parkia decussata in the upper portion (positive values), and of Acacia mangium Swietenia macrophylla Table 3. Mean annual increment and current annual increment of the diameter at and in the lower portion (negative values) (Figure 2). Overall the PCA explained 94.27% 1.30 m (MAI-DBH and CAI-DBH, respectively) and total height (MAI-Ht and CAI-Ht, respectively) of trees of five forest species, in 4-year-old homogeneous plantations in Itacoatiara, State of Amazonas, Brazil. Numbers are averages of three replicates. Each replicate was a 128 m2 plot planted with 32 seedlings.

MAI-DBH CAI-DBH MAI-Ht CAI-Ht Species (cm yr-1) (cm yr-1) (m yr-1) (m yr-1) Acacia mangium 2.1 1.2 3.3 5.4 Dipteryx odorata 1.3 1.4 1.8 2.1 Jacaranda copaia 2.7 2.6 2.8 4.3 Parkia decussata 2.4 3.0 1.7 2.4 Swietenia macrophylla 1.8 2.1 1.3 1.4

Table 4. Commercial form factor (FF), cylinder volume per area (Vcyl), mean annual increment (MAI-Vcyl) and current annual increment (CAI-Vcyl) of cylinder volume per area of four native and one exotic tree species, in 4-year-old homogeneous plantations, in Itacoatiara, State of Amazonas, Brazil. Numbers are averages of three replicates. Each replicate was a 128 m2 plot planted with 32 seedlings. Different letters following the numbers in a column indicate significant differences by the LSD test (p < 0.05).

V MAI-V CAI-V Species FF cyl cyl cil (m3 ha-1) (m3 ha-1 yr-1) (m3 ha-1 yr-1) Acacia mangium 0.51 207.48 a 51.87 120.05 Dipteryx odorata 0.50 25.39 d 6.35 15.67 Jacaranda copaia 0.42 93.57 b 23.39 57.49 Figure 1. Hierarchical clustering dendrogram for the mean of the dendrometric variables DBH (diameter at breast height, 1.30 m), CPA (crown projection area), H Parkia decussata 0.41 57.56 c 14.39 37.58 t (total height) and V (commercial cylinder volume) of 4-year-old Acacia mangium, Swietenia cyl 0.43 42.97 d 10.74 27.27 Dipteryx Odorata, Jacaranda copaia, Parkia decussata and Swietenia macrophylla macrophylla trees in monospecific forest plantations in Itacoatiara, State of Amazonas, Brazil.

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of variance in the data. Dipteryx odorata and Jacaranda copaia were both correlated to the values of Ca2+, Mg2+, pH, and sum of bases, whereas Acacia mangium and Swietenia macrophylla had correlation with the value of Al3+. We did not observe correlation of Parkia decussata with any soil chemical attributes.

DISCUSSION We observed high survival rates (> 80%) in the majority of the forest monospecific plantations at 4-year-old trees, which indicates a good adaptation to the conditions of the planting site (Nunes et al. 2015). The exception occurred for Jacaranda copaia, which had a survival rate below 70%. Apart from that, this species had the best performance results among all tested species, including both stem and crown quality, phytosanitary

and nutritional status, DBH, CPA, Ht, and Vcyl. Figure 2. Principal component analysis for the mean of some biometric variables In comparison with other studies conducted in the of trees (CC: circular crown; SPS: satisfactory phytosanitary status; GNS: good Amazon region, our survival rate of Acacia mangium was nutritional status), and soil chemical attributes at the depth of 0.00-0.30 m practically the same as observed in 1-year-old plantations, but (provided by Machado et al. 2017), at 4-year-old Acacia mangium, Dipteryx odorata, lower than in other 4-year-old plantations, and quite higher Jacaranda copaia, Parkia decussata and Swietenia macrophylla monospecific forest plantations in Itacoatiara, State of Amazonas, Brazil. pH; N; Ca2+; Mg2+; CEC: potential than 6- and 11-year-old plantations (Table 5). In contrast, the cation-exchange capacity at pH 7.0; SB: sum of bases; SOC: soil organic carbon; survival rate of Jacaranda copaia was lower in the present study SOM: soil organic carbon. than those observed in older plantations (Table 5).

Table 5. Review of seedling performance of Acacia mangium, Dipteryx odorata, Jacaranda copaia, Parkia decussata and Swietenia macrophylla in the

Amazon region. Numbers for survival rate (S), total height (Ht), diameter at 1.30 m (DBH), and cylinder volume per area (Vcyl) are mean values

Spacing Age S H DBH Vcyl Species/source t m years % m cm m3 ha-1 Acacia mangium Miranda and Valentim (2000) 1.5 x 1.5 1 80.0 2.87 - - Souza et al. (2004) 3 x 2 4 100.0 14.0 9.5 181.3 Tonini et al. (2006) 3 x 2 6 25.6 11.4 12.3 76.8 Souza et al. (2008) 3 x 3 11 27.8 - - - Souza et al. (2010) 3 x 3 6 30.6 14.5 24.4 - This study 2 x 2 4 81.0 13.3 8.5 207.5 Dipteryx odorata Souza et al. (2008) 3 x 3 11 80.6 10.4 9.6 48.6 Souza et al. (2010) 3 x 3 6 86.1 8.1 6.6 - This study 2 x 2 4 99.0 7.1 5.2 25.4 Jacaranda copaia Sampaio et al. (1989) 3 x 4 9 63.0 13.47 18.83 12.7 4.5 x 4 9 79.0 19.43 23.53 10.3 Tonini et al. (2006) 4 x 3 5 73.4 7.6 10.1 17.9 This study 2 x 2 4 68.0 11.3 10.9 93.67 Parkia decussata This study 2 x 2 4 95.0 6.6 9.8 57.6 Swietenia macrophylla Souza et al. (2008) 3 x 3 11 36.1 12.0 16.1 136.4 Souza et al. (2010) 3 x 3 6 50.0 7.7 10.9 - Escalante et al. (2012) 5 x 5 1 26.9 (site 1); 78.0 (site 2) 3.9 (site 1); 8.1 (site 2) - - This study 2 x 2 4 98.0 5.4 7.1 42.9

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Acacia mangium presented at least four times more stem We expected to observe an increase in CPA in Swietenia bifurcations and three times more tortuous stems than the macrophylla, because of the super growth of lateral buds native species in our study, which was also observed by Tonini caused by the attack of all of the individuals by Hypsipylla et al. (2006). Both characteristics are desirable for biomass grandella. The appearance of numerous lateral branches production. Acacia mangium has a tendency to form multiple reduces significantly the commercial value of the wood trunks, thus the individuals of this species presented the of this species (Souza et al. 2008). Thus, the attack by the highest number of stems (units per hectare), while the other insects compromised the nutritional status of the trees, studied species had lower and similar values. Tonini et al. which inhibited the vigorous growth of several shoots and (2006) also observed a 45.7% stem bifurcation in Jacaranda also contributed to the low growth in height and diameter of copaia, while we had practically no stem bifurcation, nor Swietenia macrophylla, in comparison to the other species. The tortuous stems, for this species in our study. Tree species with strong smell of young trees of Swietenia macrophylla attracts rectilinear stems are indicated for the production of this moth, that lays its eggs on the shoots, usually on the dorsal and laminates (Trevisan et al. 2013). surface of leaves, originating caterpillars that cause serious Apart from plantation age, the variability in performance damages to the apical meristem, delaying growth, and even within species observed in Table 5 may also have been leading to the death of affected plants after successive attacks influenced by differences in other parameters, such as spacing, (Lunz et al. 2009). Attacks of H. grandella contributed to the soil characteristics, climatic conditions, quality of seedlings, low survival (36.1%) of Swietenia macrophylla in plantations et al and preparation of the planting area (Barbosa et al. 2003). In in Manaus (Souza . 2008). Therefore, monospecific plantations of Swietenia macrophylla should be avoided and 16-year-old monospecific plantations of Jacaranda copaia in the planting of this species should only occur if measures are Manaus (Amazonas State) the survival rates of trees, as well as taken to minimize this problem, such as intercropping with DBH, increased with increased spacing (Sampaio et al. 1989). Jacaranda copaia, Parkia decussata or Dipteryx odorata, since Despite their comparatively low rates of stem bifurcation the diversification of forest species and agroforestry systems (< 20%), Dipteryx odorata and Swietenia macrophylla probably considerably reduce infestation in Swietenia macrophylla require the removal of the less vigorous branch of the trees (Silva et al. 2013b). The overwhelming satisfactory bifurcation to obtain straight and more vigorous stems. These phytosanitary and nutritional status of the other three native species can be used for furniture production and finishing species indicated that they had higher resistance to pests and articles, since they have noble wood with easy workability pathogens, which is a relevant information about the scarcely (Almeida et al. 2010; Zau et al. 2014). known nutritional requirements of native forest species of the The competition among plants may also affect seedling Amazon biome (Santos Filho et al. 2007). survival. In Bolivia, survival rates in two 1-year-old plantations Our height values of Acacia mangium were similar, (5 x 5 m spacing) of Swietenia macrophylla were 26.9% and whereas the values of DBH and Vcyl were lower, compared to 78%, and were likely related to a higher herbaceous and those obtained for older plantations (Table 5). Four-year-old cover in the former plantation site (Escalante et al. 2012), plantations of Acacia mangium with increased spacing (3 x 2 since Swietenia macrophylla is a gap-demanding species (Dias m) had higher values of DBH (9.5 cm) (Souza et al. 2004) and Marenco 2006). than in this study, because a wider spacing positively affects The large CPA observed in Dipteryx odorata, Parkia the diameter growth of trees (Nascimento et al. 2012). Souza decussata and Jacaranda copaia is a desirable characteristic et al. (2008) did not evaluate the growth of Acacia mangium in forest plantations, since the tree crown is responsible in 11-year-old plantations because the survival rate was too for reducing the direct impact of raindrops and the direct low (< 30%), and recommended that this species should incidence of solar radiation on the soil (Ignácio et al. 2007). be harvested between 3 and 5 years of age in monospecific Therefore, a rapid soil cover contributes to the stabilization plantations, when their survival rate was 66.7 % and 36.1 of soil aggregates and the reduction of weed-control costs. %, respectively. Their results probably were influenced by the Besides, circular crowns, as is the case with Jacaranda copaia, occurrence of shoot and stem breaks, and windblow falls of are considered an interesting silvicultural feature, because they trees higher than 6 m. have uniform growth, thus favoring the planning of more The values of both height and DHB ofDipteryx odorata, efficient spacing between trees (Condéet al. 2013). The low Jacaranda copaia and Swietenia macrophylla in our study values of CPA for Acacia mangium and Swietenia macrophylla tended to be higher than those of older plantations (Table were probably related to high competition among the trees, 5). In 11-year-old monospecific plantations of ten native and due to the high number of stems per hectare (5138 stem ha- exotic forest species in Manaus, the lowest values of height, 1). Among Acacia mangium trees, 14% had signs of fungal DBH and cylinder volume occurred for Dipteryx odorata infection, which may hinder the growth of this species in (Souza et al. 2008). This pattern is probably a characteristic plantations (Halfeld-Vieira and Nechet 2010). of this species, which grows mainly in primary forests and

15 VOL. 48(1) 2018: 10 - 17 ACTA AMAZONICA MACHADO et al. Silvicultural performance of forest species has high wood density, one of the most resistant among performance, because of the position that this species occupied Amazonian timber species (Stangerlin et al. 2013). The in the principal component analysis. DBH growth of Jacaranda copaia did not differ significantly among different spacings (Sampaioet al. 1989). The species CONCLUSIONS also responded with higher DBH values when the soil was Jacaranda copaia Dipteryx odorata Parkia harrowed (Barbosa et al. 2003). Successful plantation results , followed by and have raised the interest in Jacaranda copaia as a silvicultural decussata, are suitable for planting in areas with edaphoclimatic species in the Amazon region (Tonini et al. 2008). conditions similar to those of the present work, due to the best performance in 4-year-old monospecific experimental We found no results for diametric growth of Parkia plantations, in comparison to Acacia mangium and Swietenia decussata seedlings in the literature. However, Salomão et macrophylla. The lowest performance ofSwietenia macrophylla al. (2014) recommended the use of this species in forest was influenced by the severe attack ofHypsipylla grandella. restoration after mining in conservation units in Amazonia, because its seedlings had 100% survival after 13 years. The good diametric growth ofParkia decussata, combined REFERENCES with its excellent debranching (Murilo Rezende Machado, ABRAF - Associação Brasileira de Produtores de Florestas Plantadas. personal observation), and low frequency of stem bifurcation 2013. Anuário estatístico ABRAF 2013 ano base 2012. ABRAF, or tortuous stems, rendered this species as suitable for the Brasília. 148p. production of lumber. In the State of Roraima, a 5-year-old Almeida, A.N.; Angelo, H.; Silva, J.C.G.L.; Hoeflich, V.A. 2010. plantation (4 x 3 m spacing) of Parkia multijuga Benth. had Mercado de madeiras tropicais: substituição na demanda de higher DBH (12.3 cm) (Tonini et al. 2006) than Parkia exportação. Acta Amazonica, 40: 119-126. decussata in the present study. Barbosa, A.P.; Campos, M.A.A.; Sampaio, P.T.B.; Nakamura, S.; The high values of both MAI-H and CAI-H for Acacia Gonçalves, C.Q.B. 2003. O crescimento de duas espécies t t florestais pioneiras, pau-de-balsa ( lagopus Sw.) e caroba mangium and Jacaranda copaia indicated that, at four years (Jacaranda copaia D. Don), usadas para recuperação de áreas of age, these species would already require a wider spacing to degradadas pela agricultura na Amazônia central, Brasil. Acta maintain and to stimulate their current annual increment in Amazonica, 33: 477-485. diameter. At least in the case of Acacia mangium, this pattern was Barros, L.S.; Vale Júnior, J.F.; Schaefer, C.E.G.R.; Mourão Júnior, corroborated by the high rate of stem bifurcation, which implies M. 2009. Perdas de solo e água em plantio de Acacia mangium intense competition for light. The rapid growth in height of Wild e savana em Roraima, norte da Amazônia. Revista Brasileira Acacia mangium, also reported by Miranda and Valentim de Ciência do Solo, 33: 447-454. (2000), may be influenced by the high capacity of this species Condé, T.M.; Lima, M.L.M.; Lima Neto, E.M.; Tonini, H. to explore the nutritional pool in the soil, which causes soil 2013. Morfometria de quatro espécies florestais em sistemas acidification due to cation depletion (Yamashitaet al. 2008). agroflorestais no munícipio de Porto Velho, Rondônia. Revista

Generally, Vcyl of Dipteryx odorata increased with the age Agroambiente, 7: 18-27. of plantations, while that of Jacaranda copaia and Swietenia Costa, K.C.P.; Fereira, M.J.; Linhares, A.C.C.; Guedes, A.V. 2015. macrophylla decreased in older plantations (Table 5). The Vcyl of Biomassa e nutrientes removidos no primeiro desbaste em plantio Acacia mangium (207.48 m3 ha-1) and Jacaranda copaia (93.57 de Bertholletia excelsa Bonpl. Scientia Forestalis, 43: 591-600. 3 -1 m ha ) were very high compared with those observed by Souza Dias, D.P.; Marenco, R.A. 2006. Photoinhibition of photosynthesis et al. (2004) in 4-year-old plantations of A. mangium (3 x 2 m in Minquartia guianensis and Swietenia macrophylla inferred by 3 -1 spacing) (181.26 m ha ) and Tonini et al. (2006) for Jacaranda monitoring the initial fluorescence.Photosynthetica , 44: 235-240. 3 copaia in 5-year-old plantations (4 x 3 m spacing) (17.98 m Diniz, M.B.; Oliveira Junior, J.N.; Trompieri Neto, N.; Diniz, M.J.T. -1 ha ). In both situations, the spacing used was higher than that 2009. Causas do desmatamento da Amazônia: uma aplicação used in the present study (2 x 2 m). Therefore, spacing clearly do teste de causalidade de Granger acerca das principais fontes influenced our results, since a smaller spacing produces higher de desmatamento nos municípios da Amazônia Legal brasileira. volumes of wood per hectare (Nascimento et al. 2012). Nova Economia, 19: 121-151. The better performance of Dipteryx odorata and Jacaranda Escalante, E.; Pando, V.; Saravia, P.; Bravo, F. 2012. Survival and growth copaia was correlated with the higher soil fertility (higher of big- (Swietenia macrophylla King) seedlings in availability of calcium and magnesium, which influenced the two provenance trials in Bolivia. Ecología en Bolivia, 47: 37-52. highest values of pH and sum of bases at the depth of 0.00- Falesi, I.C.; Cruz; E.S; Pereira, F.B.; Lopes, E.C. 1969. Os solos da área 0.30 m; Machado et al. 2017). On the other hand, the worst de Manaus-Itacoatiara. Instituto de Pesquisa e Experimentação performance of Acacia mangium and Swietenia macrophylla Agropecuária do Norte (IPEAN), Belém. 116 p. was influenced by higher values of exchangeable aluminum Galeão, R.G; Carvalho, J.O.P.; Yared, J.A.G.; Marques, L.C.T.; Costa in the soil, which can be toxic and compromise the growth Filho, P.P. 2006. Diagnóstico dos projetos de reposição florestal and survival of most plants. Parkia decussata had intermediate no Estado do Pará. Revista de Ciências Agrárias, 45: 101-120.

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Comportamento Polímeros, 24: 726-732. nutricional de espécies arbóreas utilizadas no reflorestamento de áreas degradadas sob o impacto da exploração petrolífera na RECEIVED: 01/03/2017 região de Urucu, Município de Coari, AM. Revista Brasileira de ACCEPTED: 20/09/2017 Biociências, 5: 1134-1136. ASSOCIATE EDITOR: Francesco Ripullone

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